, Volume 43, Issue 1, pp 1–16 | Cite as

Diel nectar secretion rhythm in squash (Cucurbita pepo) and its relation with pollinator activity

  • Andrea A. Edge
  • Byron N. van Nest
  • Jennifer N. Johnson
  • Samara N. Miller
  • Nick Naeger
  • Sam D. Boyd
  • Darrell MooreEmail author
Original article


Most studies of foraging behavior in bees have been performed under artificial conditions. One highly neglected area is the daily nectar secretion rhythm in flowers including how nectar properties may vary with time of day. As a first step in understanding the connections between forager behavior and nectar presentation under more natural conditions, we examined nectar secretion patterns in flowers of the squash Cucurbita pepo. Under greenhouse conditions, squash flowers exhibit consistent diel changes in nectar volume and concentration through anthesis. These temporal patterns are robust, persisting under field conditions as well as simulated drought conditions in the greenhouse. In the presence of active pollinators, diel patterns are evident but with highly variable, severely reduced volumes. The potential consequences of these factors for pollinator behavior are discussed.


bees foraging nectar pollinators behavior 



We thank Huijing Sun, Caleb Paquette, Curt Gill, Joanna Magner, and Aprele Fitzgerald for valuable assistance with the greenhouse and 2008 field experiments. We very much appreciate the generosity of the ETSU Kingsport Campus for providing garden space and support for the 2008 field studies and Thomas Jones and Allison Mains for garden space and support at their farm in 2009. Financial support was provided by a grant from the United States Department of Agriculture, National Research Initiative, CSREES #2006-35302-17278 (DM).


  1. Beier, W. (1968) Beeinflussung der inneren Uhr der Bienen durch Phasenverschiebung des Licht-Dunkel-Zeitgebers. Z Bienenforschung 9, 356–378Google Scholar
  2. Beier, W., Lindauer, M. (1970) Der Sonnenstand als Zeitgeber für die Biene. Apidologie 1, 5–28CrossRefGoogle Scholar
  3. Beling, I. (1929) Über das Zeitgedächtnis der Bienen. Z. Vergl. Physiol. 9, 259–338CrossRefGoogle Scholar
  4. Boch, R. (1956) Die Tänze der Bienen bei nahen und fernen Trachtquellen. Z. Vergl. Physiol. 38, 136–167CrossRefGoogle Scholar
  5. Bolten, A.B., Feinsinger, P., Baker, H.G., Baker, I. (1979) On the calculation of sugar concentration in flower nectar. Oecologia 41, 301–304CrossRefGoogle Scholar
  6. Butler, C.G. (1945) The influence of various physical and biological factors of the environment on honeybee activity: an examination of the relationship between activity and nectar concentration and abundance. J. Exp. Biol. 21, 5–12Google Scholar
  7. Canto-Aguilar, M.A., Parra-Tabla, V. (2000) Importance of conserving alternative pollinators: assessing the pollination efficiency of the squash bee, Peponapis limitaris in Cucurbita moschata. J. Insect Conserv. 4, 203–210CrossRefGoogle Scholar
  8. Corbet, S.A. (1978) Bee visits and the nectar of Echium vulgare L. and Sinapis alba L. Ecol. Entomol. 3, 25–37CrossRefGoogle Scholar
  9. Corbet, S.A., Delfosse, E.S. (1984) Honeybees and the nectar of Echium plantagiueum L. in southeastern Australia. Aust. J. Ecol. 9, 125–139CrossRefGoogle Scholar
  10. Corbet, S.A., Wilmer, P.G. (1981) The nectar of Justicia and Columnea: composition and concentration in a humid, tropical climate. Oecologia 51, 412–418CrossRefGoogle Scholar
  11. Corbet, S.A., Willmer, P.G., Beament, J.W.L., Unwin, D.M., Prys-Jones, O.E. (1979) Post-secretory determinants of sugar concentration in nectar. Plant Cell Environ. 2, 293–308CrossRefGoogle Scholar
  12. Frisch, B., Aschoff, J. (1987) Circadian rhythms in honeybees: entrainment by feeding cycles. Physiol. Entomol. 12, 41–49CrossRefGoogle Scholar
  13. Giurfa, M., Núñez, J.A. (1992) Foraging by honeybees on Carduus acanthoides: pattern and efficiency. Ecol. Entomol. 17, 326–330CrossRefGoogle Scholar
  14. Greggers, U., Kuettner, A., Mauelshagen, J., Menzel, R. (1993) Optimization of honeybees with balanced US-qualities. In: Elsner, N., Heisenberg, M. (eds.) Gene–Brain–Behavior, Proceedings of the 21st Göttingen Neurobiology Conference, p. 841. Georg Thieme, StuttgartGoogle Scholar
  15. Kearns, C.A., Inouye, D.W. (1993) Techniques for pollination biologists. University Press of Colorado, NiwotGoogle Scholar
  16. Kleber, E. (1935) Hat das Zeitgedächtnis der Bienen biologische Bedeutung? Z. Vergl. Physiol. 22, 221–262CrossRefGoogle Scholar
  17. Lindauer, M. (1948) Über die Einwirkung von Duft- und Geschmacksstoffen sowie underer Faktoren auf die Tänze der Bienen. Z. Vergl. Physiol. 31, 348–412CrossRefGoogle Scholar
  18. Linnaeus, C. (1751) Philosophia botanica. Kiesewetter, StockholmGoogle Scholar
  19. Louw, G.N., Nicolson, S.W. (1983) Thermal, energetic, and nutritional considerations in foraging and reproduction of the carpenter bee Xylocopa capitata. J. Ent. Soc. S. Africa 46, 227–240Google Scholar
  20. Moore, D., Doherty, P. (2009) Acquisition of a time-memory in forager honey bees. J. Comp. Physiol. A 195, 741–751CrossRefGoogle Scholar
  21. Moore, D., Rankin, M.A. (1983) Diurnal changes in the accuracy of the honeybee foraging rhythm. Biol. Bull. 164, 471–482CrossRefGoogle Scholar
  22. Moore, D., Siegfried, D., Wilson, R., Rankin, M.A. (1989) The influence of time of day on the foraging behavior of the honeybee, Apis mellifera. J. Biol. Rhythms 4, 305–325PubMedCrossRefGoogle Scholar
  23. Naeger, N., Van Nest, B.N., Johnson, J.N., Boyd, S.D., Southey, B.R., Rodriguez-Zas, S.L., Moore, D., Robinson, G.E. (2011) Neurogenomic signatures of spatiotemporal memories in time-trained forager honey bees. J. Exp. Biol. 214, 979–987PubMedCrossRefGoogle Scholar
  24. Nepi, M., Guarnieri, M., Pacini, E. (2001) Nectar secretion, reabsorption, and sugar composition in male and female flowers of Cucurbita pepo. Int. J. Plant Sci. 162, 353–365CrossRefGoogle Scholar
  25. Nieh, J.C., Contrera, F.A.L., Ramirez, S., Imperatriz-Fonseca, V.L. (2003) Variation in the ability to communicate three-dimensional resource location by stingless bees from different habitats. Anim. Behav. 66, 1129–1139CrossRefGoogle Scholar
  26. Nuñez, J.A. (1966) Quantitative Beziehungen zwischen den Eigenschaften von Futterquellen und den Verhalten von Sammelbienen. Z. Vergl. Physiol. 53, 142–164Google Scholar
  27. Nuñez, J.A. (1977) Nectar flow by melliferous flora and gathering flow by Apis mellifera ligustica. J. Insect Physiol. 23, 265–275CrossRefGoogle Scholar
  28. Parker, R.L. (1925) The collection and utilization of pollen by the honeybee. Cornell University Press, IthacaGoogle Scholar
  29. Proctor, M., Yeo, P., Lack, A. (1996) The natural history of pollination. HarperCollins, LondonGoogle Scholar
  30. Rabinowitch, H.D., Fahn, A., Meir, T., Lensky, Y. (1993) Flower and nectar attributes of pepper (Capsicum annuum L.) plants in relation to their attractiveness to honeybees (Apis mellifera L.). Ann. Appl. Biol. 123, 221–232CrossRefGoogle Scholar
  31. Renner, M. (1955) Über die Haltung von Bienen in geschlossenen, künstlich beleuchteten Räumen. Naturwissenschaften 42, 539–540CrossRefGoogle Scholar
  32. Schua, L. (1952) Untersuchungen über den Einfluss meteorlogischer Elemente auf das Verhalten der Honigbiene (Apis mellifica). Z. Vergl. Physiol. 34, 258–277CrossRefGoogle Scholar
  33. Seefeldt, S., De Marco, R.J. (2008) The response of the honeybee dance to uncertain rewards. J. Exp. Biol. 211, 3392–3400PubMedCrossRefGoogle Scholar
  34. Seeley, T.D. (1986) Social foraging by honeybees: how colonies allocate foragers among patches of flowers. Behav. Ecol. Sociobiol. 19, 343–354CrossRefGoogle Scholar
  35. Seeley, T.D. (1989) Social foraging in honeybees: how nectar foragers assess their colony's nutritional status. Behav. Ecol. Sociobiol. 24, 181–199CrossRefGoogle Scholar
  36. Seeley, T.D. (1995) The wisdom of the hive. Harvard University Press, CambridgeGoogle Scholar
  37. Seeley, T.D., Camazine, S., Sneyd, J. (1991) Collective decision-making in honey bees: how colonies choose among nectar sources. Behav. Ecol. Sociobiol. 28, 277–290CrossRefGoogle Scholar
  38. Silva, E.M., Dean, B.B. (2000) Effect of nectar composition and nectar concentration on honey bee (Hymenoptera: Apidae) visitations to hybrid onion flowers. J. Econ. Entomol. 93, 1215–1221CrossRefGoogle Scholar
  39. Tepedino, V.J. (1981) The pollination efficiency of the squash bee (Peponapis pruinosa) and the honey bee on summer squash (Cucurbita pepo). J. Kansas Entomol. Soc. 54, 359–377Google Scholar
  40. Vansell, G.H. (1934) Relation between the nectar concentration in fruit blossoms and the visits of honeybees. J. Econ. Entomol. 2, 943–945Google Scholar
  41. Varju, D., Nuñez, J.A. (1991) What do foraging honeybees optimize? J. Comp. Physiol. A 169, 729–736CrossRefGoogle Scholar
  42. Vidal, M.D., De Jong, D., Wien, H.C., Morse, R.A. (2006) Nectar and pollen production in pumpkin (Cucurbita pepo L.). Revista Brasil. Bot. 2, 267–273Google Scholar
  43. Visscher, P.K., Seeley, T.D. (1982) Foraging strategy of honeybee colonies in a temperate deciduous forest. Ecology 63, 1790–1801CrossRefGoogle Scholar
  44. von Buttel-Reepen, H.B. (1900) Sind die Bienen Reflexmaschinen? Biol. Zbl. 20, 1–82Google Scholar
  45. von Frisch, K. (1942) Die Werbetänze der Bienen und ihre Auslösung. Naturwissenschaften 30, 269–277CrossRefGoogle Scholar
  46. von Frisch, K. (1967) The dance language and orientation of bees. Harvard University Press, CambridgeGoogle Scholar
  47. Waddington, K.D. (1982) Honeybee foraging profitability and round dance correlates. J. Comp. Physiol. 148, 279–301CrossRefGoogle Scholar
  48. Waddington, K.D. (1985) Cost–intake information used in foraging. J. Insect Physiol. 31, 891–897CrossRefGoogle Scholar
  49. Wahl, O. (1932) Neue Untersuchungen über das Zeitgedächtnis der Bienen. Z. Vergl. Physiol. 16, 529–589Google Scholar
  50. Wahl, O. (1933) Beitrag zur Frage des biologischen Bedeutung des Zeitgedächtnisses der Bienen. Z. Vergl. Physiol. 18, 709–717Google Scholar
  51. Wainselboim, A.J., Roces, F., Farina, W.M. (2002) Honeybees assess changes in nectar flow within a single foraging bout. Anim. Behav. 63, 1–6CrossRefGoogle Scholar
  52. Waller, G.D. (1972) Evaluating responses of honey bees to sugar solutions using an artificial-flower feeder. Ann. Ent. Soc. Am. 65, 857–862Google Scholar
  53. Wolf, S., Lensky, Y., Paldi, N. (1999) Genetic variability in flower attractiveness to honeybees (Apis mellifera L.) with the genus Citrullus. HortScience 34, 860–863Google Scholar
  54. Wyatt, R., Broyles, S.B., Derda, G.S. (1992) Environmental influences on nectar production in milkweeds (Asclepias syriaca and A. exaltata). Am. J. Bot. 79, 636–642CrossRefGoogle Scholar

Copyright information

© INRA, DIB-AGIB and Springer Science+Business Media B.V. 2011

Authors and Affiliations

  • Andrea A. Edge
    • 1
  • Byron N. van Nest
    • 1
  • Jennifer N. Johnson
    • 1
  • Samara N. Miller
    • 1
  • Nick Naeger
    • 2
  • Sam D. Boyd
    • 1
  • Darrell Moore
    • 1
    Email author
  1. 1.Department of Biological SciencesEast Tennessee State UniversityJohnson CityUSA
  2. 2.Department of EntomologyUniversity of Illinois at Urbana-ChampaignUrbanaUSA

Personalised recommendations